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Body Size and Age of the China Wood Frog (Rana Chensinensis) in Northeastern China

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Body Size and Age of the China Wood Frog (Rana Chensinensis) in Northeastern China NORTH-WESTERN JOURNAL OF ZOOLOGY 7 (2): 236-242 ©NwjZ, Oradea, Romania, 2011 Article No.: 111129 www.herp-or.uv.ro/nwjz Body size and age of the China Wood Frog (Rana chensinensis) in northeastern China Bing Yao CHEN1,3, Wen Bo LIAO2,3,* and Zhi Ping MI2,3 1. College of Life Science, Nanjing Normal University, Nanjing, 210097, China. 2. Key Laboratory of Southwest China Wildlife Resources Conservation Ministry of Education, China West Normal University, Nanchong, 637009, China. 3. Institute of Rare Animals and Plants, China West Normal University, Nanchong 637009, China. *Correspondence at: W.B. Liao, E-mail: [email protected]. Received: 03. January 2011 / Accepted: 28. July 2011 / Available online: 09. August 2011 Abstract. Age structure and body size of a population of the China Wood Frog (Rana chensinensis) in northeastern China were determined by using skeletochronology. Lines of arrested growth (LAGs) recorded in phalanges were used to estimate the age of adults. Results showed that average age did not differ significantly between males and females. Age at sexual maturity in both sexes was 1 yr. The maximum observed longevity was 5 yrs in males and 4 yrs in females, respectively. Average body size of females was significantly larger than males. A non-significant negative correlation between body size and air temperature was found among the seven populations when the effect of altitude, latitude and age was controlled, suggesting that latitudinal variation in body size of Rana chensinensis across different temperature environment do not follow Bergmann's rule. Key words: skeletochronology, Bergmann's rule, Rana chensinensis, body size, age structure. Introduction morphosed juveniles first emerge about 2 months after egg-laying (Lu et al. 2006). The body size and Bergmann's rule states that body size of endo- age and growth rate of this species are strongly in- therms tend to be larger at high latitudes or high fluenced by ambient temperature across small ele- altitudes (Bergmann 1847). This trend holds for vation ranges in Shanxi province, northern China the intraspecific body size variation in birds and (Ma 1985, Lu et al. 2006, Ma et al. 2009a). mammals (Freckleton & Pagel 2003, Meiri & Skeletochronology is a reliable method of age Dayan 2003). Body size variation of ectotherms estimation by counting the lines of arrested has the same trend (Ray 1960). However, for ecto- growth (LAGs) recorded in long bones in am- therms such as amphibians, there is considerable phibians (Hemelaar & Van Gelder 1980) and, has controversy about the direction of latitudinal been successfully used in assessing age for desert clines at the intra- and interspecific level (Ashton zones (Sullivan & Fernandez 1999), tropical re- 2002, Laugen et al. 2005, Olalla-Tarraga & Rodri- gions (Khonsue et al. 2000, Kumbar & Pancharatna guez 2007, Adams & Church, 2008, Cvetkovic et 2001), subtropical regions (Morrison et al. 2004, al. 2009). In most anurans, age and body size of Lai et al. 2005, Liao & Lu 2010a,b,c, Liao et al. 2010, individuals at high altitudes or high latitudes Liao & Lu 2011a, Li et al. 2010) and temperate re- (cold temperature) being older and larger than at gions (Guarino et al. 2003, Guarino & Erismis low altitudes or low latitudes (i.e. warm tempera- 2008, Kyriakopoulou- Sklavounou et al. 2008, Ma ture; Ashton 2002, Morrison et al. 2004, Lai et al. et al. 2009b, Üzüm 2009, Sinsch et al. 2010). 2005, Lu et al. 2006, Olalla-Tarraga & Rodriguez Here, we carried out a study for body size and 2007, Liao & Lu 2010b,c, Liao et al. 2011, Liao & Lu age in R. chensinensis using the data from a popu- 2011b). lation in northeastern China where these demo- The China Wood Frog (Rana chensinensis) is graphic parameters are unavailable. Moreover, we native to the montane regions of northern China used data on body size and age from six R. (Tanaka-Ueno et al. 1999). This frog is typically re- chensinensis populations based on the previous stricted to montane rivers, where it is the domi- studies (Lu et al. 2006, Ma et al. 2009a). Our aims nant anuran (Liu and Hu 1961, Xie et al. 2000). were to compare age, body size, and growth traits Rana chensinensis has a relatively long hibernating between the sexes in the population in northeast- period (3-5 months). Egg-laying lasts for 3-4 ern China and analyze variation in age, body size, weeks, and the frog may be classified as an explo- and growth among seven populations with differ- sive spring breeder (Wells 1977). Newly meta- ent temperature. Size and age of Rana chensinensis 237 Materials and Methods Kolmogorov–Smirnov test to identify age structure be- tween the sexes. Multifactorial analyses of covariance The fieldwork was conducted from January to February (ANCOVA) were used to assess the difference in body 2010 at Caohezhang Town (124º01′E, 41º04′N, and eleva- size between sexes when the effects of age were removed. tion 367 m) in Benxi County of Liaoning Province in The relationship between body size and age was analyzed northeastern China. The area has annual average tem- using one linear regression. We used partial correlation perature of 6-8°C (sub-zero mean monthly temperatures analysis to test variation in body size among populations occur in January–February) and annual total precipitation across latitudes with altitude and age as covariables. All of 700-1000 mm. The vegetation covering the study site is values given are shown as mean ± SD, and statistical tests characterized by tree species Xylosma racemosum, Fraxinus were two-tailed. mandshurica, Juglans mandshurica, Pinus koraiensis, Larix gmelinii. Individuals were captured by hand during the day in Results hollows under the riverbank, or in deep ponds at their hi- bernation sites. Each individual was sexed by the secon- A total of 123 individuals (63 males and 60 fe- dary sexual traits (the nuptial pads on their thumbs in males) were captured by chance in this population adult males and of the ova in adult females). The snout– vent length (SVL) was measured using a vernier caliper to resulting in a sex ratio of 1.05: 1. Lines of arrested the nearest 0.1 mm. The longest phalange of third toe of growth (LAGs) were present in cross sections of the right hind limb of each individual was removed by phalanges for 122 adult individuals of this species toe-clipping methods, and stored in 10% neutrally buff- (Fig. 1). The endosteal resorption of LAGs incom- ered formalin at room temperature for histological sec- pletely eroded the first (innermost) periosteal. tions. LAGs were observed in 28 males and 32 females. The standard skeletochronological procedure was However, the first LAGs of 6 males and 7 females used to determine age (e.g. Liao & Lu 2010a,b, Liao et al. 2010). For each individual digit, we cleaned off surround- were completely eroded by endosteal resorption ing tissues of the phalanges, and then put them in 5% ni- due to the absence of KL, and we added one year tric acid to decalcify for 48 h. These decalcified digits as their true age. For all samplings, endosteal re- were stained for 180 min in Harris’s haematoxylin. The sorption of LAGs did not affect LAGs counted. We stained bones were dehydrated by running the bone did not find double lines and false lines in any through successive ethanol stages of 70, 80, 95, and 100% samplings. for approximately 1 h in each concentration. Tissues were Average body size differed significantly be- infiltrated with successive paraffin changes for 1 h in an ± oven at 50 °C, embedded in small paraffin blocks and al- tween males and females (males, 60.6 2.9 mm; lowed to harden overnight. Cross-sections (13 μm thick- females, 67.7 ± 4.6 mm; Student’s t – tests: t = ness) of the phalanx with the smallest medullar cavity 10.17, p < 0.001). Within each age group, females and the thickest cortical bone were selected and mounted had significant larger body size than males (Table on glass slides. The sections were observed though a light 1). microscope and the best of them were photographed with a Motic BA300 digital camera mounted on a Moticam2006 Table 1. Body size (SVL, mm) within each age class of light microscope at ×400 magnification. The analysis of the China Wood Frog (Rana chensinensis) in northeast- lines of arrested growth (LAGs) was performed by two ern China. Values in descending order are mean ± SD persons (W.B. Liao & Z.P. Mi) with previous experience and sample size. of the technique and the results were then compared for age determination until consensus was reached. The in- Age class Females Males Z P nermost LAG can be reabsorbed with growth of endosteal 1y 66.1 ± 3.7 59.0 ± 2.5 3.62 < 0.001 tissue in amphibians which would affect the accuracy of (n = 8) (n = 13) age estimates (Smirina 1983). However, previous studies 2y 66.9 ± 3.5 60.0 ± 2.2 5.96 < 0.001 have shown that the endosteal resorption of LAGs occurs (n = 26) (n = 32) in frogs primarily before they reach sexual maturity and 3y 69.0 ± 4.5 62.5 ± 3.4 3.34 0.001 slows down thereafter (Hemelaar 1985, Castanet et al. (n = 16) (n = 12) 1996, Liao & Lu 2010b, Liao et al. 2010b). Moreover, a 4y 69.0 ± 7.5 63.6 ± 1.9 2.07 0.04 weak reabsorption of LAGs in R. chensinensis was ob- (n = 9) (n = 5) served by noting the presence of the Kastschenko Line 5y 65.08 (KL; the interface between the endosteal and periosteal (n = 1) zones, Lu et al. 2006, Ma et al.
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